Conifers

Conifers are the largest, most widespread, and most economically important group of gymnosperms (nonflowering seed plants), including about 630 species divided into six or seven families. Conifers are the oldest extant group of seed plants, dating back to more than 280 million years in the fossil record. Some of the current families and genera have long fossil records; for example, remarkably well-preserved and modern-appearing cones of the genus Araucaria dating to 160 million years ago have been discovered, and a well-preserved fossil pine cone dating to 130 million years ago can be compared directly with cones of living pine trees.

Conifer Diversity

All conifers are woody plants, mostly trees or sometimes shrubs. Typical conifers such as members of the pine, cypress, and araucaria families are recognized by their woody seed cones, with flattened or shield-shape cone scales arranged spirally or in pairs or whorls around a central axis. The woody-coned conifers usually have winged seeds that are dispersed by wind and gravity. Other important groups of conifers such as the yew family, junipers, and most of the podocarp family have their seed cones reduced to one- or few-seeded fleshy structures that are dispersed by birds. Conifer seed cones range from less than 1 centimeter in length to up to 50 centimeters long and may be quite massive in some species of pines and araucarias.

Most conifers are evergreen, but a few genera (notably bald cypress, dawn redwood, and larch) shed their leaves during the winter. The majority of conifers have narrow, needle-shaped leaves, arranged in spirals or sometimes in pairs, or are found in tightly clustered whorls on short branches. Pines are unusual in having their leaves extremely tightly clustered in needle clusters (fascicles) with almost no stem elongation between the leaves. Some conifers have their leaves very reduced and scalelike (most of the cypress family), while subtropical to tropical conifers in the podocarp and araucaria families may have the leaves flattened and are relatively broad.

Conifers include some of the longest-living, tallest, and most massive trees in the world. Bristlecone pines from the southwestern United States are among the longest living individual trees in the world, having been dated from tree rings to more than five thousand years in age. Sequoias are among the tallest trees in the world, reaching more than 110 meters in height, while the related giant sequoia reaches 106 meters in height and up to 11 meters in diameter. Large and ancient specimens of conifers are featured attractions

SELECTED CONIFER GENERA
Common Name	Generic Name	Number of Species (approximate)	Family of Species	Geographic Range	Economic Uses
Pine	Pinus	Pine	110	Northern Hemisphere	Timber, paper, resins, ornamentals
Spruce	Picea	Pine	40	Northern Hemisphere	Timber, paper, ornamentals
Fir	Abies	Pine	50	Northern Hemisphere	Timber, paper, resins, ornamentals
Hemlock	Tsuga	Pine	10	Eastern and western North America, eastern Asia	Timber, paper, ornamentals
Douglas-fir	Pseudotsuga	Pine	6	Western North America, eastern Asia	Timber, paper
Juniper	Juniperus	Cypress	50	Northern Hemisphere	Wood, pencils, flavorings, ornamentals
Cypress	Cupressus	Cypress	13	Western North America, Eurasia	Ornamentals
Bald cypress	Taxodium	Cypress	2	Eastern United States to Central America	Timber, ornamentals
Sequoia	Sequoia	Cypress	1	California to southern Oregon	Timber, ornamentals
Yew	Taxus	Yew	10	North America, Eurasia	Ornamentals, medicinal alkaloids
Araucaria	Araucaria	Araucaria	18	South America, South Pacific	Timber, ornamentals
Podocarpus	Podocarpus	Podocarp	95	Southern Hemisphere, northern to eastern Asia, Mexico, Caribbean	Timber, ornamentals

in national parks in many parts of the world, most notably sequoias and giant sequoias in California, alerce (Fitzroya) in Chile and Argentina, and kauri (Agathis) in New Zealand.

Conifer Distribution

Conifers are important forest components in many areas of the world, and members of the pine family are especially abundant in cool to cold-temperate and mountainous areas of the Northern Hemisphere, where such genera as pines, spruces, firs, and hemlocks often form dense forests. Junipers and pines are also very abundant trees in semiarid environments of the Northern Hemisphere such as the Great Basin region of the western United States. Pines are the most widely distributed genus of trees in the Northern Hemisphere and are also especially widely planted as timber trees in both hemispheres.

Several genera of conifers with only a single living species of very restricted distribution were once much more widespread in the fossil record and have been termed "living fossils." These include the dawn redwood (Metasequoia) from China and the sequoia and giant sequoia from California. Another remarkable genus, Wollemia (from the araucaria family), was known only as a fossil from Australia until 1994, when a living plant of this species was discovered growing in a remote canyon area near Sydney, Australia.

Economic Uses

Conifers are extremely important economically as sources of lumber and other wood products, and are also widely planted as ornamental trees and shrubs. The most important sources of softwood lumber in the world are trees in the pine family, especially species of pine, spruce, larch, and Douglas-fir, which are widely used for dimensional timber for building construction and boat building, and for general construction uses such as utility poles, doors, and cabinetry. These woods are also widely used for plywood and veneer and as sources of wood pulp for paper and cardboard and other modified wood products, such as charcoal. Southern yellow pines, such as slash pine and loblolly pine, are widely grown in their native southeastern United States as sources of lumber and pulp, while the Monterey pine from coastal California is now widely planted as a commercial timber tree in the Southern Hemisphere. Douglas-fir (Pseudotsuga menziesii) is a particularly important timber species in the northwestern United States and Canada. Several species of pines are tapped or cut and steam-distilled for stem resins, which are used as commercial sources of turpentine, tar oils, rosin, and pitch. Wood of the Norway spruce and white spruce has also been prized for constructing musical instruments such as violins, and the light, strong wood of Sitka spruce has been used for aircraft construction. The attractive reddish-colored wood from species of the cypress family, such as the sequoia, is quite weather- and decay-resistant and is highly prized for building construction, decks, fences, and other outdoor uses. Wood of the western red cedar (Thuja plicata) has been heavily used for weather-resistant roof shingles. Fragrant wood from junipers has natural insect-repellent properties and is used for moth-resistant cedar closets or chests. Wood of juniper and incense cedar has been commonly used to make pencils.

Many species of conifers are grown as ornamentals, and a wide variety of cultivated shrub forms have been selected for garden use from members of the yew and cypress families, including several species of yew, juniper, cypress, and golden cypress (Chamaecyparis). Conifers from a number of genera are prized as ornamental trees, of which some particularly attractive examples are the blue spruce (Picea pungens), the Himalayan cedar (Cedrus deodara), and the Norfolk Island pine (Araucaria heterophylla). Several species of firs and pines are commercially grown and cut as Christmas trees, and young plants of the subtropical Norfolk Island pine are grown for indoor use as living Christmas trees. Several species of pines from Eurasia and North America are highly esteemed as sources of oil-rich edible seeds (pignoli or pine nuts). Cones of Juniperus communis (juniper berries) are used as flavorings in cooking and provide the aromatic flavoring of gin, whose name is derived through the Dutch jenever from the name of juniper. Recently, bark and leaves of several species of yews have become important as the source of taxol and related alkaloids, which disrupt the process of cell division and are used in the treatment of several types of cancer.

Robert A. Price

Bibliography

Dallimore, W., A. Bruce Jackson, and S. G. Harrison. A Handbook of Coniferae and Ginkgoaceae, 3rd ed. New York: St. Martin's Press, 1967.

Judd, Walter S., Christopher S. Campbell, Elizabeth A. Kellogg, and Peter F. Stevens. Plant Systematics: A Phylogenetic Approach. Sunderland, MA: Sinauer, 1999.

Richardson, David M., ed. Ecology and Biogeography of Pinus. New York: Cambridge University Press, 1998.

Rushforth, Keith D. Conifers. London: Christopher Helm, 1987.